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by Dr. Lena Struwe 
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updated: 01/19/11 

Anatomy and other less visible characters of gentians
(Palynology, phytochemistry, karyology)

Gentians are very variable in their anatomy and other characters. Listed below are some traits that occur in the family and their occurrence in investigated genera. Bibliographic references are listed in parenthesis [...] after the characters.

See  also the morphology page.


General anatomy [Perrot, 1897]:




latex: absent in all gentians, except from the fruits of Fagraea


Wood anatomy [Carlquist, 1986; Carlquist & Grant, 2005; Mennega, 1980; Perrot, 1897]:


vestured pits: common in gentians [Jansen & Smets, 1998, 2000]


nodal anatomy [Post, 1958]


Flower and fruit anatomy (see also Morphology) [Lindsey, 1940; Padmanabhan et al., 1978; Perrot, 1897; Struwe et al., 1997]:

corolla with fibers: Coutoubea, Lisianthius, Neurotheca [Woodson, 1938]

  orbicules (minute granules of sporopollenin on secretory tapetum cells in anthers): found in 38 species of 28 genera, in all tribes [Vinckier & Smets, 2003]

Corona at the base of the stamens: Symbolanthus only


postgenital fusion of carpels, at least in upper part of ovary: common in gentians, e.g., Centaurium, Enicostema, Gentiana, Saccifolium [Maguire & Pires, 1978]


congenital fusion of carpels: Anthocleista, Aripuana, Fagraea, Potalia



Palynology [Ho et al., 1994; Nilsson, 1968, 1970, 2002; Punt, 1978, 1980; Punt & Nienhuis, 1976]
The largest collection of microscopic slides made from Gentianaceae pollen is housed in Stockholm, Sweden, at the Palynological Laboratory of the Swedish Museum of Natural History.



monads: all gentians except those listed under tetrads and polyads.  Helieae genera with monads include Chorisepalum, Macrocarpaea, Tachia, and Zonanthus


tetrads: Potalia maguieorium, all in subtribe Coutoubeinae, and many genera in the Helieae, e.g., Adenolisianthus, Chelonanthus spp., Calolisianthus, Helia, Lagenanthus, Lehmanniella, Purdieanthus, Rogersonanthus, Symbolanthus, Tetrapollinia


polyads: Irlbachia, Chelonanthus spp., Calolisianthus spp.,


symmetrical pollen grains: all gentians except Voyria spp.


asymmetric pollen grains, reniform, bilateral: Voyria spp.


3-colporate: most gentians


(2–)3(–6)-porate: Anthocleista, Potalia, and Voyria


ora with lateral extensions: common


reticulate exine: common among gentians, especially in tribe Helieae


striato-reticulate exine: common among gentians


striate exine: many gentians, especially in tribe Gentianeae


rugulate exine:


foveolate exine: 


smooth exine:


scabrate exine: 


perforate exine:


imperforate exine:


loops on exine: 


spines on exine: Irlbachia, Tetrapollinia


verrucae on exine:


granules or globules on exine: 


Seed anatomy (see also Seeds) [Bouman et al., 2002; Grothe & Maas, 1984]:


very reduced seeds, dust seeds: Voyria spp.

  flattened, winged seeds: Chorisepalum

testa cells star- or puzzle-shaped: Exacum spp., Ornichia, Tachiadenus


Embryology [Bouman et al., 2002; Maheswari Devi, 1962; Padmanabhan et al., 1978]:


tenuinucellate: all investigated gentians


Chromosome numbers:


tetraploids: some groups in Centaurium and Sabatia


chromosome numbers: known numbers for each genus are listed in Struwe et al. (2002)


Phytochemistry [Jensen & Schripsema et al., 2002]:


seco-iridoids: present in all investigated gentians


xanthones: present in all investigated gentians from tribe Chironieae, Gentianeae, Helieae, and Potalieae [Carpenter et al., 1969; Meszaros, 1994; Rezende & Gottlieb, 1973]


See morphology page for Morphology
spp. = some species


References and publications

Bisset, N. G. 1980a. Phytochemistry. Pp. 211-233. In: A. J. M. Leeuwenberg, editor. Engler and Prantl’s Die natürlichen Pflanzenfamilien, Angiospermae: Ordnung Gentianales, Fam. Loganiaceae, vol. 28b (1). Duncker and Humblot, Berlin.

Bouman, F., L. Cobb, N. Devente, V. Goethals, P. J. M. Maas, & E. Smets. 2002. The seeds of Gentianaceae. Pp. 498-572. In: L. Struwe & V. A. Albert, editors. Gentianaceae – systematics and natural history. Cambridge University Press, Cambridge.

Carpenter, I., H. D. Locksley, & F. Scheinmann. 1969. Xanthones in higher plants: biogenetic proposals and a chemotaxonomic survey. Phytochemistry 8: 2013-2026.

Carlquist, S. & J. R. Grant. 2005. Wood anatomy of Gentianaceae, tribe Helieae, in relation to ecology, habit, systematics, and sample diameter. Brittonia 57(3): 276-291.

Daniel, M. & S. D. Sabnis. 1978. Chemical systematics of family Gentianaceae. Curr. Sci. 47: 109-111.

Grothe, E. H. M. & P. J. M. Maas. 1984. A scanning electron microscopic study of the seed coat structure of Curtia Chamisso & Schlechtendahl and Hockinia Gardner (Gentianaceae). Proc. Kon. Ned. Akad. Wetensch., Ser. C, 87 (1): 33-42.

Jansen, S. & E. Smets. 1998. Vestured pits in some woody Gentianaceae. IAWA Journal 19: 35-42.

Jansen, S. & E. Smets. 2000. Morphology, distribution, and systematic importance of vestures in the Gentianales. In: Nordenstam, B., El-Ghazaly, G. & M. Kassas (eds.) Plant systematics for the 21st century: 277-296, 25 figs. Wenner-Gren International Series Volume 77, Portland Press, London. XIII-366 p. ISBN 1 85578 135 2

Jensen, S. R. & J. Shripsema. 2002. Chemotaxonomy and pharmacology of Gentianaceae. Pp. 573- 632. In: L. Struwe & V. A. Albert, editors. Gentianaceae – systematics and natural history. Cambridge University Press, Cambridge.

Maguire, B. & J. M. Pires. 1978. Saccifoliaceae – a new monotypic family of the Gentianales. Pp. 230-245. In: B. Maguire & collaborators, editors. The Botany of the Guayana Highland – Part X. Mem. New York Bot. Gard. 29. 

Marloth, R. 1909. A diplostigmatic plant, Sebaea exacoides (L.) Schinz (Belmontia cordata L.). Trans. Roy. Soc. South Africa 1(1): 311-314.

Meszáros, S. 1994. Evolutionary significance of xanthones in Gentianaceae: a reappraisal. Biochem. Syst. Ecol. 22: 85-94.

Nilsson, S. 1964. On the pollen morphology in Lomatogonium A. Br. Grana Palynol. 5: 298-329.

Nilsson, S. 1967a. Pollen morphological studies in the Gentianaceae-Gentianinae. Grana Palynol. 7: 46-145.

Nilsson, S. 1967b. Notes on pollen morphological variation in Gentianaceae-Gentianinae. Pollen Spores 9: 49-58.

Nilsson, S. 1968. Pollen morphology in the genus Macrocarpaea (Gentianaceae) and its taxonomical significance. Sv. Bot. Tidskrift 62: 338-364.

Nilsson, S. 1970. Pollen morphological contributions to the taxonomy of Lisianthus L. s. lat. (Gentianaceae). Sv. Bot. Tidskrift 64: 1-43.

Nilsson, S. 2002. Gentianaceae – a review of palynology. Pp. 377-497. In: L. Struwe & V. A. Albert, editors. Gentianaceae – systematics and natural history. Cambridge University Press, Cambridge.

Nilsson, S. & J. J. Skvarla. 1969. Pollen morphology of saprophytic taxa in the

Padmanabhan, D., D. Regupathy, & S. Pushpa Veni. 1978. Gynoecial ontogeny in Enicostemma littorale Blume. Proc. Indian Acad. Sci. 87 B (Pl. Sci.-2, no. 5): 83-92. 

Perrot, M. E. 1897 [1899]. Anatomie comparée des Gentianacés. Ann. Sci. Nat. Bot. Ser. 8, 7: 105-292.

Post, D. M. 1958. Studies in Gentianaceae. I. Nodal anatomy of Frasera and Swertia perennis. Botanical Gazette. 120(1):1-14.

Punt, W. 1978. Evolutionary trends in the Potalieae (Loganiaceae). Rev. Palaeobot. Palynol. 26: 313-335. [Anthocleista, Fagraea, Potalia]

Punt, W. 1980. Pollen morphology. Pp. 162-191. In: A. J. M. Leeuwenberg, editor. Engler and Prantl’s Die natürlichen Pflanzenfamilien, ANGIOSPERMAE: Ordnung Gentianales, Fam. Loganiaceae, vol. 28b(1). Duncker and Humblot, Berlin.  [Anthocleista, Fagraea, Potalia]

Punt, W. & P. W. Leenhouts. 1967. Pollen morphology and taxonomy in the Loganiaceae. Grana Palynol. 7: 469-516. [Anthocleista, Fagraea, Potalia]

Punt, W. & W. Nienhuis. 1976. The northwest European pollen flora. 6. Gentianaceae. Rev. Palaeobot. Palynol. 21: NEPF 89-123.

Rezende, C. M. A. D. M. & O. R. Gottlieb. 1973. Xanthones as systematic markers. Biochem. Syst. 1: 111-118.

Struwe, L., J. W. Kadereit, J. Klackenberg, S. Nilsson, M. Thiv, K. B. von Hagen, & V. A. Albert. 2002. Systematics, character evolution, and biogeography of Gentianaceae, including a new tribal and subtribal classification. Pp. 21-309. In: L. Struwe & V. A. Albert (eds.), Gentianaceae: Systematics and Natural History, Cambridge University Press, Cambridge.

Vinckier, S. & E. Smets. 2003. Morphological and ultrastructural diversity of orbicules in Gentianaceae. Annals of Botany 92: 657-672.

Woodson, R. E. 1938. Observations on the floral fibres of certain Gentianaceae. Ann. Bot. (London) 50: 759-766.

© Lena Struwe, 2002-2004.


© Gentian Research Network, 2002-2011.
For corrections and additions, contact Lena Struwe at